The Fossil Flamingos of Australia
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July, 1963 289 THE FOSSIL FLAMINGOS OF AUSTRALIA By ALDEN H. MILLER When the abundant bird fossils collected in recent years in the Lake Eyre basin of Australia were first tentatively assorted, the most surprising discovery was that of flamingos (Stirton, Tedford, and Miller, 1961:35). This type of bird is absent from the continent of Australia today and flamingos do not occur at points nearer than north- west India; there are records in historic times from Ceylon, and their bones occur along with remains of the Dodo in Mauritius. The further study of our collections, especially those obtained in 1961, shows that flamingos were present not only in mid-Tertiary time as previously noted but also in the Pleistocene, although strangely they did not appear among the Pleistocene collections DeVis ( 1906) reported on from Cooper Creek in the Lake Eyre area. In all, there are four species represented, three of which are here described as new. In two of the faunas two kinds of flamingo occur together. This is not an unusual situation in this order of birds. Three specieslive today in the high- land playa lakes of the arid sections of Chile, and the presence of several species to- gether in the Oligocene of France was well documented by Milne-Edwards (1867-1871). Flamingos in general are well represented in the fossil record, both the typical group, the family Phoenicopteridae, and the extinct Paloelodidae, a group of birds that is straight-billed and also less slim and elongate than the true flamingos. The environ- ment in which such birds live, namely relatively shallow, muddy lakes of wide expanse, has been particularly favorable to the preservation of their bones. Thus the record of the order extends back to the Cretaceous and geographically to many parts of Europe and North America. Fossils and modern occurrences are not known from Southeast Asia or the’East Indies. The lack of fossil records for South America and Africa prob- ably reflects in part the.relatively slight attention given to the fossil birds of those continents. The support of the National Science Foundation in 1961 and 1962 made possible further field work in Australia and the processing of collections, both of the essential Recent bird skeletons and of fossils. The study of fossil vertebrates in South Australia has continued to receive the generous support of the South Australian Museum and its staff. We were especially aided in 1961 by Norman B. Tindale, Paul F. Lawson, and Harry J. Bowshall. R. A. Stirton, Richard H. Tedford, and Virginia D. Miller of our field party provided critical assistance during the 1961 expedition. Stirton particularly has given constant support and encouragement to the investigation of the Australian fossil birds and has supplied details concerning the collecting localities. I am indebted also to Philip S. Humphrey of the United States National Museum for the loan of skeletons of Phoenicopterus ruber. DESCRIPTION OF MATERIAL Phoenicopterus novaehollandiae, new species Type.-Right tarsometatarsus, the distal end essentially complete, although with many frac- tures proximal to the articular surfaces; about 95 per cent of shaft represented; no. P13648, South Australian Mus.; locality no. V6150, Univ. Calif. Mus. Paleo., Lake Pitikanta; Ngapakaldi Fauna, middle Tertiary, probably late Oligocene or early Miocene; figures la, c, 2a. Type locality.-Etadunna Formation in escarpment on west side Lake Pitikanta about 600 yards from south end; Lake Eyre region, South Australia; green to greenish-brown mudstone below upper gray calcareous mudstone and near contact with lower gray calcareous mudstone. 290 THE CONDOR Vol. 65 Diagnosis.-Similar in foot structure to Phoenicopterus ruber (includes Phoenicopterus ruber rosew) but scar for articulation of metatarsal I present; distal tarsal foramen apparently not per- forating plantar surface and on anterior surface situated more proximally; trochlea IV closer to trochlea III; posterior extension of trochlea II narrower mediolaterally and proximodistally. Size within limits of variation of ruber but near maximum of range. Analysis and comparison.-The species novaehollandiae has the same general con- figuration, slender construction and articular system of the foot as that of the modern greater flamingo, P. ruber, of Africa, southern Europe, and the West Indies. The most interesting departure is in the presence of the articulation for a first toe which can only mean that this digit was of a size and functional importance much greater than in modern flamingos. In the present-day Phoenicopterus it is reduced to a length of about 18 mm. and seems to have little use. In the genus Phoeniconaias it is further reduced and in Phoenicopawus it is lacking. Even in the first two, there is no flattened scar for articulation at the point of attachment on the medial ridge of the plantar surface, the toe being held at that point rather loosely by weak ligaments. :I j a b C d Fig. 1. Distal ends of right tarsometatarsi of flamingos, natural size. a, type of Phoenicopterw novaehollandiae, medial view; b, modern Phoenicopterus ruber roseus, no. 289137, U. S. Nat. Mus., medial view; c, type of P. novaehollandiae, plantar view ; d, P. Y. yoseus, plantar view. Drawings in figures 1 to 6 by Augusta Lucas. The other features of Phoenicopterus novaehollandiae in respect to the trochleae reflect a lesser spread of the toes and perhaps weaker support for the medial bracing ligaments of the inner toe. These differences are of small magnitude and probably lim- ited functional significance but are consistent departures from the configuration in the July, 1963 FOSSIL FLAMINGOS OF AUSTRALIA modern ruber in that none of the considerable sam- ple of rubier shows them. They are therefore useful speciescharacters. Phoenicopterus novaehollandiae had an overall length and slenderness equivalent to specimen no. 289737, U. S. Nat. Mus., a large male P. ruber (figs. lb, d, 2b). This length is judged from the dis- tance from the distal end to the beginning of the expansion of the proximal end, which expansion is present in the fossil even though the proximal artic- ular surfaces and the hypotarsus are entirely lack- ing. The shaft, which was lying in place, but in a much fragmented condition, has been fitted together, all junctions but one being natural contacts. Because of the one gap in mid-shaft, which is believed to be small, the estimate of length is minimal, but it is not likely to be more than 5 per cent below the true value. Five extinct species of flamingos of the genus Phoenicopterus have already been described from other parts of the world. Two of these are very small species,neither of which is represented by the distal end of the tarsometatarsus. By reason of small size at least, they are not confusable with novaehollan- diae. These two are Phoenicopterus minutus Howard (19.55) from the late Pleistocene of California and Phoenicopterus stock L. Miller (1944) from the Pliocene of Chihuahua. Phoenicopterus copei Shufeldt ( 1892 ) of the Ple- istocene of Oregon, consisting of several unassociated parts, does include a distal segment of a tarsometa- tarsus as designated type material. Howard (1946: 157-158) did not find characteristics in this part of the tarsus either in size or shape which separated copei from ruber, in the inclusive sense, although there were features of the tibiotarsus that did so dis- tinguish it. The figure of the tarsometatarsus pre- pared by Shufeldt himself shows this bone in an- terior aspect to have none of the compression of the trochleae of novaehollandiae, and a completely per- forate distal foramen is present and situated as in ruber. b Phoenicopterus floridanus Brodkorb (1953) from the Pliocene of Florida is based on a tibiotarsus but there are referred specimens of the tarsometatarsus. Brodkorb’s figure of this element shows none of the characteristic shape of trochlea II of novaehollan- Fig. 2. Anterior views of tarsometa- tarsi, X %. a, type of Phoenicop- diae. Moreover the shaft is much deeper in floridanus terus novaehollandiae ; b. modern and the distal foramen is perforated and situated Phoenicopterus ruber roseus, no. more as in ruber. 289737, U.S. Nat. Mus. 292 THE CONDOR Vol. 65 The only old world fossil of the genus Phoenicoptems which has been given a specific name is Phoenicopterus croizeti Gervais of the Aquitanian (Oligocene) of France. This was founded on a tibiotarsus. Much additional material has been referred to it, including several tarsometatarsi. Milne-Edwards (1867-1871 :pl. 81) in figuring these tarsi shows none of the features that would suggestnovaehollandiae. For example trochlea II is not compressedor tapered posteriorly and the lateral rim of trochlea IV is not produced and therefore does not reflect compression; in both these respects it is like ruber. The distal foramen clearly is placed rather far distally, not proximally as in novaehollandiae. Unfortunately the plantar surface is not figured or commented on so that evidence for the development of digit I cannot be assessed.It would be most interesting to know if croizeti as a species of similar age to novaehollandiae gave indi- cation of having a similar large hind toe unlike the later flamingos. TABLE 1 MEASUREMENTS IN MILLIMETERS OF TARSOMETATARSI OF SPECIES OF Phoenicopterus Total Le$$;;:h Le$&w;h W;;Ah.agss length P. novoehollundioe 3542 6.9 7.0 17.6 ’ P. ruber roseus, no. 289737 ( $ ) U.S.N.M.’ 356 6.2 6.9 21.6 P. ruber roseus, no. 224858 U5N.M.l 364 5.6 6.7 20.6 P. ruber roseus, no. 79026 M.V.Z. 275 5.8 6.4 18.5 P. ruber ruber, no. 140923 M.V.Z. (Galapagos Islands) 253 5.7 6.2 18.0 P. chilensis, no. 125159 ($ ) M.V.Z. 276 5.6 5.3 19.3 P.